The classification of athyridid brachiopods

Abstract

Phylogenetic relationships among higher taxa of articulate brachiopods that developed a spiralium as mineralized support for the lophophore, and the resulting classifications, are not universally agreed upon due to the complex pattern of character evolution within the phylum. Our results, using phylogenetic methodology in conjunction with traditional methods, showed a more complete and easily testable picture of this evolution than either method used alone. A detailed working hypothesis of athyridide phylogeny and the supporting evidence on character distribution is given, and a new classification of the impunctate and punctate athyrids sensu lato is presented with diagnoses to subfamily. Agreement between the stratigraphical first appearance of athyridid subfamilies and their cladistic rank was quite good (except for the koninckinoids), suggesting that both outgroup and traditional paleontological methods indicate a similar direction of character polarity in the evolution of the group. The classification is proposed based on a mix of external and internal characters which persisted during unbroken lineages. New taxa are the families Parazygidae and Triathyrididae; the subfamilies Whitfieldellinae, Triathyridinae, Septathyridinae, Cleiothyridininae, Ochotathyridinae and Plectospirinae, and the genera Brimethyris, Carteridina, Tetraloba and Cooperispira. We studied the great morphological diversity among the athyridids. Their complete stratigraphical ranges help in building phyletic lineages.